Two viruses, one a nuclear polyhedrosis virus and the other a granulosis virus, were applied in an attempt to initiate epizootics in populations of western spruce budworm, Choristoneura occidentalis Freeman, on Douglas-fir trees, Pseudotsuga menziesii (Mirb.) Franco, in southeastern British Columbia. Two 172-ha plots were aerially treated in 1982 with 9.0 kg of lyophilized, virus-infected larval powder that was formulated in an emulsifiable oil tank mix and applied at 9.4 L per hectare. Each plot was treated when larval populations were at the peak of the fourth instar. The nuclear polyhedrosis virus was applied at 5.4 × 1011 polyhedral inclusion bodies per hectare and the granulosis virus at 1.7 × 1014 capsules per hectare. Results showed that the granulosis virus treatment caused 34.6% population reduction (Abbott’s formula) and the nuclear polyhedrosis virus 51.8%. Larvae from treated and check plots were reared individually in the laboratory and the incidence of viruses, parasitoids, and successful adult emergence was recorded. Studies m these plots continued in 1983 and 1984. Although vertical transmission of both viruses was evident, their impact on budworm mortality was less than in 1982. Consequently, the epizootics were not sufficiently intense to control the target insect population.

We carried out a 2-year study to elucidate the biology of the gregarious, idiobiont ectoparasitoid Elachertus cacoeciae (Howard) by placing (implanting) laboratory-reared spruce budworm larvae [Choristoneura fumiferana (Clemens)] on current-year balsam fir (Abies balsamea L.) shoots in the field, simulating low (endemic) densities of the budworm. Spring female E. cacoeciae attacked fourth-, fifth-, and sixth-instar budworm larvae, beginning near the predicted peak of the fourth instar and ending about 10–12 days after the predicted peak of the pupal stage of the wild budworm population. The mean (±SE) brood size of spring females was 2.9 ± 0.3 E. cacoeciae pupae per host. The proportion of females increased during the season, with many broods consisting of 100% females late in the season. In 1994 and 1995, the mean proportion of females was 0.74 ± 0.05 and 0.79 ± 0.05, respectively. In the laboratory, development time from eggs to adults was approximately 20 days at 20.6 °C. Adult males provided with honey water lived 43.6 ± 3.2 days, whereas females provided with hosts and honey water lived 90.1 ± 6.6 days. Spring females had a pre-oviposition period of 11.5 ± 1.3 days, resulting in a generation time (egg to egg) of ~31 days. The oviposition period lasted 76.3 ± 7.7 days during which time spring females parasitized 19.2 ± 1.9 hosts, and produced a clutch size of 4.9 ± 0.4 eggs per host, for a lifetime fecundity of 96.8 ± 14.7 eggs. The post-oviposition period was 18.5 ± 3.7 days. Throughout their lifetime, spring females host fed only (host feeding without oviposition) on an additional 9.3 ± 1.9 hosts. Approximately 2% of pupae developing from spring females overwintered, whereas approximately 95% of pupae developing from summer females overwintered. Laboratory results for summer females suggest that they may be adapted to parasitizing alternate host(s) rather than spruce budworm.

Nutritional variability in resources may cause differential mortality between sexes resulting in biased sex ratios. If males and females differ in fitness, then mortality of the more sensitive sex can cause a bias in sex ratios, and can stimulate dispersion of males. We reared three generations of spruce budworm (Choristoneura fumiferana (Clemens); Lepidoptera: Tortricidae) on two artificial diets: a “normal” diet that provided all nutritional requirements for development and a “stress” diet (deficient in sugars and slightly higher in nitrogen), that simulated deterioration of food quality during outbreak conditions and had a detrimental impact on larval survival, development and growth. We tested the effects of continued nutritional stress on the sex ratio of pupae and adults. We found biased sex ratios in favour of males related to diet. Low quality food resulted in fewer females. This distortion was observed from the second generation onward, with a lower percentage of females reaching the pupal and adult stage. These results provide evidence that nutritional variation causes differential mortality between sexes, suggesting that females are more sensitive to nutritional stress. This is the first study that demonstrates sex ratio distortion due to nutritional selection pressure in spruce budworm. Our results indicate the importance of studying sex ratio distortion of spruce budworm in outbreak conditions.

Collections (68) of spruce budworm from 33 locations from Newfoundland to Alaska were analysed for isozyme frequencies using horizontal starch gels. Collections represented pre-, early-, mid-, late-, and post-outbreak stages of several populations in balsam fir, white spruce, and mixed host forests, as well as successive annual collections at several locations. Isozymes were measured at 11 loci in mature larvae and at six loci in pheromone-trapped males; frequencies were essentially the same in both stages, and from all host species. Three loci (IDH-2, LDH-1, and AAT-1) were found to be sex-linked, with no heterozygotes in females. Mean percentage heterozygosity ranged from 13.2 to 23.1; at individual locations it tended to decrease over successive years of outbreak and over successive collections in the same year. Contingency chi-square analysis indicated small differences related to location and outbreak history but all populations were generally homogeneous over the entire range. Nevertheless, one allozyme of AAT-1 exhibited a significant cline in frequency from the southeast to the northwest. Gene flow across the entire range appeared to be appreciable.

A standardized bioassay procedure was used to determine median lethal doses (LD50) of the microsporidium, Nosema fumiferanae (Thom.), on newly molted fourth- and fifth-instar eastern spruce budworm larvae (Choristoneura fumiferana (Clem.)). The LD50 for fifth-instar larva was 1.23 × 106 ± 2.82 × 105 spores. The fourth-instar LD50 was 2.23 × 104 ± 4.30 × 103 spores per larva for populations experiencing prolonged post-diapause cold storage or an elevated temperature during diapause and 2.00 × 105 ± 6.66 × 104 spores per larva for populations not experiencing stressful conditions during and after diapause. Median lethal times (LT50) ranged from 6 to 19 days, depending on instar and dose level. Sublethal responses of fourth- and fifth-instar larvae inoculated with serial dilutions of spores were estimated by significant linear models. These regressions were negative for pupal weight and adult longevity and positive for development time (duration of instar VI). Inoculations of newly molted sixth-instar larvae produced similar models, although development time was not significantly affected. Insects reared following stress during and after diapause had consistently longer developmental times. The importance of prolonged developmental time on disease expression and insect susceptibility is discussed.

Glypta fumifermae (Vier.) (Ichneumonidae) and Apanteles fumiferanae Vier. (Braconidae) are two of the most common parasites of immature larvae of the spruce budworm, Choristoneura fumiferana (Clem.). Females parasitize the minute host larvae in the fall. Their eggs or larvae lie dormant within the hibernating host larvae, and then resume development in the spring, when the spruce budworm larva breaks hibernation and commences to feed. The budworm host is killed by Apanteles usually when it is in the fifth larval instar or by the Glypta when it is in the sixth instar.

Aerial application of Mimic® 2LV to rising outbreak populations of the spruce budworm (Choristoneura fumiferana (Clemens); Lepidoptera: Tortricidae) in Québec, Canada, resulted in high levels of population reduction at spray deposits of 0.5–1.2 μg tebufenozide/g of foliage. Application to potted host trees in outdoor enclosures followed by bioassays revealed multiple effects on spruce budworm survival and recruitment. Chronic (14-day) exposure of late-instars to treated foliage reduced larval survival and also pupal survival, mating success, and fecundity, depending on the product concentration applied. Treatments that produced foliar deposits of ~ 0.5–1.5 μg tebufenozide/g caused high larval mortality. Exposure to deposits of ~ 0.15–0.5 μg/g caused delayed mortality during the pupal stage and reduced the mating success of survivors, while exposure to ~ 0.07–0.15 μg/g reduced the fecundity of mated females. Sublethal exposure did not affect the progeny of survivors, either in egg hatch, survival during diapause, or survival and performance after diapause. Reduced survival during late-larval and pupal stages combined with lower recruitment as a result of reduced mating success and fecundity are likely to play a role in the suppression of Mimic®-treated spruce budworm populations in the years following treatment.

The insecticide phosphamidon, both alone and mixed with other chemicals, was tested in the laboratory as a motor stimulant to female spruce budworm moths. For phosphamidon alone, the time to reaction of virgin female budworm (Choristoneura fumiferana) moths was about 40 min. However, small supplements of pyrethrum or synthetic pyrethroids reduced the reaction time to about 3–8 min. The slow reaction to phosphamidon alone could account for the failure of that insecticide to kill gravid females. It is postulated that moths quickly stimulated to flight activity within the canopy would be more likely to acquire a lethal dose of insecticide before the spray cloud dissipated.

The eastern spruce budwonn, Choristoneuva fumiferana (Clem.) (Lepidoptera: Tortricidae), is subject to a variety of naturally occurring infectious diseases including nuclear polyhedrosis virus (NPV), granulosis virus (GV), cytoplasmic virus (CPV), and entomopox virus (EPV), of which the most intensively studied is NPV (CfMNPV) (Cunningham 1985). If CfMNPV is ever to be deemed an effective and economical alternative to chemical pesticides for spruce budworm control, additional research is required.

The role of adult nutrition in longevity, progeny production, and offspring sex ratio of Trichogramma minutum Riley was examined. On average, honey-fed females lived 26.4 days and produced 260 offspring; unfed females lived 3.5 days and produced 80 offspring. Feeding on fructose or sucrose also significantly increased longevity and fecundity over unfed females (fructose, 23 days and 230 offspring; sucrose, 21 days and 230 offspring) but to a lesser degree than feeding on pure honey. Females fed yeast suspension or water had no significant increases in longevity or fecundity compared to unfed females. Offspring sex ratios of long-lived females were male-biased (50–62% males), those of short-lived females were female-biased (74–82% females). Lifetime reproduction of honey-fed females was highest at 20–25 °C and relative humidities of 20–80%, but short-term offspring production (during the first 2 days after emergence) was highest at 30 °C and 60–80% RH. Females that had access to honey for only a 24-h period did not increase their offspring production over the first 4 days of their lives when compared to unfed females. The potential benefits of feeding adult T. minutum for mass-rearing and field release are discussed.

Allozymes at several polymorphic loci were assayed in larval collections of 12 recognized species and two possible new species of Choristoneura and two species of Archips. Most of the 48 collections came from high density populations, and those of C. fumiferana, C. occidentalis, and C. pinus represented much of the geographic range of these species. Mean percentage heterozygosity ranged from 2.0 to 18.6%, based on nine polymorphic loci. Three loci are sex-linked in C. fumiferana, two in C. pinus and C. occidentalis and probably in some other members of the group. Allozymes of aspartate transaminase (AAT-1) were most varied among the species and permit identification of individual C. fumiferana in better than 95% of cases. Among the group of coniferophagous Choristoneura species genetic distances were small (max. Nei = 0.232); C. fumiferana was the most distinct species. Wagner trees based on modified Rogers’ distances supported the above conclusions but indicated that separations among C. biennis, C. orae, C. occidentalis, C. carnana, C. subretiniana, and the two new species of Choristoneura were very small and probably below the species level, based on the allozymes measured.

During 1982 and 1984, ground releases of Trichogramma minutum Riley were assessed for control of the spruce budworm, Choristoneura fumiferana (Clemens), on 12- to 20-year-old, white spruce stands in northern Ontario. Maximum parasitism of susceptible egg masses was 16 and 87% following the release of 480 000 and 12 million female T. minutum per hectare, respectively. Releases at intervals of 1 week maintained parasitism of susceptible egg masses at constant levels throughout the oviposition period of spruce budworm. When parasitism of susceptible egg masses was maintained above 78.2% during the ovipositional period, total egg mass parasitism averaged 58.0% and resulted in an 80.3% reduction of overwintering 2nd-instar larvae. The optimal strategy for reducing spruce budworm was two releases of T. minutum at an interval of 1 week in the ovipositional period. This allowed a second generation of parasitoids to emerge from the spruce budworm eggs that were more efficient in maintaining high levels of parasitism than those emerging from the standard rearing host. Natural parasitism of spruce budworm egg masses was less than 4% and there was no carryover of parasitism in the years following inundative release. The rate of T. minutum release necessary to achieve effective mortality of spruce budworm during outbreak populations is discussed briefly.

Annual spruce budworm, Choristoneura fumiferana (Clemens), survey data were subjected to multiple and logistic regression analyses to examine the relationship between egg mass density in the fall and resultant defoliation the next season. Egg mass density was the most important variable associated with resultant defoliation, followed by current defoliation, regional population trends, host species, and sprays. Together, these accounted for 60% of the variation in resultant defoliation. Balsam fir [Abies balsamea (L.) Miller] suffered greater levels of defoliation than white spruce [Picea glauca (Moench) Voss] at a given egg mass density. Resultant defoliation of balsam fir also showed a steeper response to egg mass density than resultant defoliation of white spruce. Levels of current defoliation increased susceptibility to defoliation in a similar manner between species, as did regional population trends. Sprays were more effective at reducing resultant defoliation on balsam fir than on white spruce but, overall, did not confer a high level of foliage protection. Predictions of resultant defoliation using the multiple regression models had confidence limits averaging 75%, which are too large to be useful for predictive purposes. The logistic regression equations could be used to predict the probability of a stand receiving light or severe defoliation.

Outbreak and declining populations of spruce budworm (Choristoneura fumiferana (Clem.)) were sampled extensively at three locations in New Brunswick, Canada, between 1982 and 1992 and were examined for the prevalence of granulosis and nuclear polyhedrosis viruses (Baculoviridae). Larvae, pupae, and adults were collected using a variety of methods. Spruce budworm nuclear polyhedrosis virus (CfMNPV) genomic DNA probes and wet-mount light microscopy were used to determine CfMNPV prevalence in 50 274 juvenile spruce budworms. Spruce budworm granulosis virus (ChfuGV) genomic DNA probes were used to determine the prevalence of ChfuGV in 25 703 of these same samples. The prevalence of both viruses was low, with ChfuGV and CfMNPV not found in more than 15% and 2%, respectively, of samples in any collection in a given year. Prevalence of ChfuGV was greatest in mid- to late June in sixth-instar larvae. Each virus was detected in only two of 2177 female moths and in none of the 420 male moths examined. In the entire collection, cytoplasmic polyhedrosis virus (Reoviridae) was detected in only two budworm larvae and entomopoxvirus (Poxviridae) was not detected in any.

Disruption of mating and of trap capture of spruce budworm, Choristoneura fumiferana (Clem.), has been accomplished in recent years (reviewed in Silk and Kuenen 1984) by disseminating large quantities of the primary components (synthetic) of the female sex pheromone within a forest environment. These components (E)- and (2)-11-tetradecenal (E/Z11-14:Ald) (Sanders and Weatherston 1976) are emitted by virgin female budworm at a 95/5 E/Z ratio (Silk et al. 1980). Unsaturated aldehydes are expensive, tend to polymerize readily (Dunkelblum et al. 1984), and are susceptible to air oxidation and/or photodegradation in the field environment. Analogues, with increased stability and reduced cost, that duplicate the effects of these pheromone components would therefore be of practical value.

The spruce coneworm, Dioryctria reniculleloides (Mut. and Mun.), is often associated with the spruce budworm on spruce host trees and sometimes approaches or equals the spruce budworm, Choristoneura fumiferana (Clem.), in numbers (Spies and Dimond 1985). The coneworn is probably~responsiblefo r some of the damage on spruce attributed to the budworm, and with much balsam fir in eastern North America now dead from defoliation or harvested, there is increasing interest in spruce as the major resource to protect.

A white spruce, Picea glauca (Moench) Voss (Pinaceae), plantation in southern Quebec was found to contain two distinct types of trees, the first resistant and the second susceptible to attack by spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae). To identify the mechanisms of white spruce resistance to spruce budworm, we studied the role of epicuticular waxes, comparing (i) the foliar chemistry of susceptible and resistant trees and (ii) the feeding pattern of larvae at first contact with the foliage. Needles collected from resistant trees contained concentrations of the monoterpenes α-pinene and myrcene that were 307% and 476%, respectively, above those found in needles collected from susceptible trees. Although there were no significant differences in probing behaviour, significantly fewer larvae transitioned from probing to feeding on resistant needles; this led to fewer feeding bouts as well as a significantly shorter first meal. Removal of waxes increased the number of individuals transitioning from probing to feeding on resistant needles; this led to more feeding bouts. Our results demonstrate that monoterpenes influence the pattern of feeding of spruce budworm larvae as well as playing an important role in white spruce resistance.

The behavioral responses of male spruce budworm moths to a wide range of loadings of synthetic sex pheromone on filter paper and rubber septa were investigated in a wind tunnel. The highest proportion of males flying upwind and reaching filter-paper dispensers occurred at pheromone loadings of between 0.1 and 10 μg. Above these loadings, males were activated but upwind flight was arrested before the moths reached the pheromone source. No such arrestment occurred with rubber septa up to the maximum loading assayed, 1 mg. Up to 72% of the males that reached a septum attempted to copulate with it, but even this response was less than that to a septum in the presence of pheromone emitted by females. This confirms previous conclusions that the synthetic pheromone blend used here, 95:5 (E:Z-11-tetradecenal) is incomplete.

The 45-cm mid-crown branch tip from balsam fir and white spruce is described in terms of surface area, fresh weight, and bud density. Fresh weight is suggested as the most appropriate unit to express density of all stages of the spruce budworm’s, Choristoneura fumiferana (Clem.), life cycle, particularly for the purposes of comparisons between host species and locations with different foliage conditions.

Changes in distribution of the spruce budworm in the crown of balsam fir are documented for all stages of the insect’s life cycle. Correction factors to account for these changes when estimating density on the basis of the 45-cm branch tip sampling unit are given for balsam fir and white spruce.

The amount of error in detection of spruce budworm larvae on foliage of both host trees by sampling personnel varied systematically and consistently as a function of insect development. A method to compensate for this type of error is also suggested.